Morphological and molecular description of a novel species of Eimeria (Apicomplexa) that infects extraintestinal tissues of kiwi (Aves: Apteryx spp.)
Syst Parasitol (2025) 102:30
https://doi.org/10.1007/s11230-025-10227-x
Morphological and molecular description of a novel species
of Eimeria (Apicomplexa) that infects extraintestinal tissues
of kiwi (Aves: Apteryx spp.)
Emma Scheltema
Laryssa Howe
· Kerri Morgan
· Preet Singh
· Barbara Adlington ·
Received: 31 January 2025 / Accepted: 23 March 2025 / Published online: 9 April 2025
© The Author(s) 2025, Corrected publication 2025
Abstract Coccidia (Apiconmplexa) are naturally
occurring and occasionally detrimental parasites of
kiwi (Apteryx spp.), a unique, flightless bird species
dependent upon conservation efforts for survival.
Using morphological and molecular data, a new coccidia species, Eimeria koka n. sp., isolated from two
closely related but geographically isolated kiwi host
species, Apteryx rowi Tennyson et al. (rowi) and
Apteryx mantelli Bartlett (North Island brown kiwi),
is described. Oocysts are oval (20.8 × 15.9 μm) with a
mean L/W ratio of 1.3, and a distinctive rough, crenellated brown oocyst wall (mean 1.2 μm), an oocyst
residuum, 1–2 polar granules, and no micropyle. Sporocysts are ellipsoidal (11.6 × 6.3 μm) with a Stieda
body and sporocyst residuum. Phylogenetic analysis
of the cytochrome C oxidase subunit 1 (CO1) placed
E. koka n. sp. in a separate clade to other Eimeria
species previously identified from kiwi (Coker et al.,
Syst Parasitol 100(3):269–281, 2023). Comparison of
DNA from oocysts with infected tissues from a single
Supplementary Information The online version
contains supplementary material available at https://doi.
org/10.1007/s11230-025-10227-x.
E. Scheltema (*) · P. Singh · B. Adlington · L. Howe
School of Veterinary Sciences, Massey University,
Palmerston North, New Zealand
e-mail:
K. Morgan
Wildbase, Massey University, Palmerston North,
New Zealand
juvenile North Island brown kiwi confirmed parasitism of the kidney and lung tissues. This is the first
Eimeria species identified from extraintestinal tissues
in kiwi. Further molecular studies are recommended
to determine the tissue distribution of E. koka n. sp.
and other Eimeria species in kiwi.
Introduction
Kiwi, the smallest extant species of ratite, are nocturnal, flightless ground-dwelling birds with an unusual burrow-nesting behaviour (Calder, 1978; Taborsky & Taborsky, 1995; Sales, 2005; Peat, 2006).
There are five recognised species of kiwi (Apterygiformes: Apterygidae) in two morphological groups,
all endemic to New Zealand: brown kiwi consisting
of North Island brown kiwi (Apteryx mantelli Bartlett), rowi (Apteryx rowi Tennyson et al.), and tokoeka
(Apteryx australis Shaw), and spotted kiwi consisting
of little-spotted (Apteryx owenii Gould) and greatspotted kiwi (Apteryx haastii Potts) (Burbidge et al.,
2003; Shepherd et al., 2012).
Kiwi populations have declined dramatically over
the last century as a result of escalating anthropogenic impacts upon the natural environment, including habitat modification and successive introductions of mammalian predators (McLennan et al.,
1996, 2004). Since the early 1990s, with the launch
of the Kiwi Recovery Plan, there has been a significant investment in conservation interventions, such
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as captive-rearing (Operation Nest Egg or ONE), to
maintain and, in some cases, increase kiwi populations (Butler & McLennan, 1991; Colbourne et al.,
2005; Robertson et al., 2011; Germano et al., 2018).
However, Coccidia (Apicomplexa), which are ubiquitous in captive and wild kiwi populations, are considered a major limiting factor in the success of captiverearing programmes, causing morbidity and mortality
in juvenile captive birds (Boardman, 2008; Morgan
et al., 2014).
Coccidiosis in kiwi was first recorded in a 19-dayold captive-reared brown kiwi chick suffering from
renal infection in 1978 (Thompson & Wright, 1978).
Four coccidia morphotypes were reported from faeces of other kiwi chicks from the same facility, but
they were unable to be sporulated or further described
(Thompson & Wright, 1978). The first in-depth epidemiological, morphological, and molecular study of
coccidia in kiwi was carried out by Morgan (2013).
This study included a thorough analysis of sporulated oocysts from North Island brown kiwi, and the
author was able to confirm that the species of coccidia routinely recovered from kiwi were of the genus
Eimeria. Further, Morgan described four distinct species; Eimeria apteryxii (Morgan et al., 2017), E. kiwii
(Morgan et al., 2017), E. paraurii (Morgan et al.,
2017), and E. mantellii (Morgan et al., 2017) from
North Island brown kiwi. Subsequently, one additional species from North Island brown kiwi, Eimeria
paopaoii (Coker et al., 2023), has been described.
Infection with multiple species of Eimeria is common in kiwi, and while infection typically occurs in
the intestine, it occasionally occurs in extraintestinal
tissues, including the kidney, liver, lungs, and spleen
(Morgan et al., 2012, 2013). However, which Eimeria
species cause these differing disease pathologies is
currently unknown.
These previous studies have described coccidia
recovered from a relatively small number of individual North Island brown kiwi from a limited number of
geographic locations. Although most Eimeria species
are highly host specific, recent research shows that
some have a wider host spectrum than traditionally
thought; several Eimeria species are shared across
multiple host genera or family groups (Duszynski
& Wilber, 1997; Mácová et al., 2018; Kvičerová
et al., 2020; Trefancová et al., 2021). Coker (2021)
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reported Eimeria species from Haast tokoeka (A.
australis “Haast”) that were morphologically synonymous with four of the species described to date from
North Island brown kiwi. Given that there has been
limited investigation of coccidia infecting other species of kiwi across a variety of geographic locations,
it is possible that more kiwi Eimeria species have
yet to be identified and described. In addition, translocations of kiwi for the purposes of conservation
inadvertently bring together hosts and parasites that
are normally geographically isolated, and may have
facilitated parasite transmission between members of
the genus Apteryx (Morgan et al., 2017; Coker, 2021;
Jahn et al., 2022).
In the present study, a morphologically distinct
species of Eimeria recovered from both rowi and
North Island brown kiwi, from multiple geographic
locations around New Zealand, is described. Establishing basic biology and taxonomy of Eimeria species in kiwi is a fundamental step in developing tools
to detect and manage disease.
Materials and methods
Morphological analysis
Faecal sample collection and storage
Coccidia-positive faecal samples were received from
routine diagnostic screening of kiwi carried out at
Massey University’s Parasitology Department (Massey University, Manawatū, New Zealand). Samples
were submitted from captive and creche (protected
wild) location (...truncated)